Editing Parasitoid (section)

==Taxonomic range==

About 10% of described insects are parasitoids, in the orders [[Hymenoptera]], [[Diptera]], [[Coleoptera]], [[Neuroptera]], [[Lepidoptera]], [[Strepsiptera]], and [[Trichoptera]]. The majority are wasps within the Hymenoptera; most of the others are Dipteran flies.<ref name=Godfray1994/><ref>{{cite journal |last1=Eggleton |first1=P. |last2=Belshaw |first2=R. |year=1992 |title=Insect parasitoids: an evolutionary overview |journal=Philosophical Transactions of the Royal Society B |volume=337  |issue=1279 |pages=1–20 |doi=10.1098/rstb.1992.0079|bibcode=1992RSPTB.337....1E }}</ref><ref>{{cite book |last1=Foottit |first1=R. G. |last2=Adler |first2=P. H. |year=2009 |title=Insect Biodiversity: Science and Society, Volume 1 |page=656 |publisher=Wiley-Blackwell |isbn=978-1118945537}}</ref> Parasitoidism has [[convergent evolution|evolved independently]] many times: once each in Hymenoptera, Strepsiptera, Neuroptera, and Trichoptera, twice in the Lepidoptera, 10 times or more in Coleoptera, and no less than 21 times among the Diptera. These are all [[holometabolous]] insects ([[Endopterygota]], which form a single [[clade]]), and it is always the larvae that are parasitoidal.<ref name="Foottit 2017"/> The metamorphosis from active larva to an adult with a different body structure [[Exaptation|permits]] the dual lifestyle of parasitic larva, freeliving adult in this group.<ref>{{cite web|title=Parasites (Parasitoids)|url=http://www.biology.ualberta.ca/courses.hp/ent207/lec31-32.htm|publisher=University of Alberta|access-date=10 March 2018|archive-date=2 March 2018|archive-url=https://web.archive.org/web/20180302202049/http://www.biology.ualberta.ca/courses.hp/ent207/lec31-32.htm|url-status=dead}}</ref> These relationships are shown on the [[phylogenetic tree]];<ref name=Niehuis2012>{{cite journal |author1=Niehuis, O. |author2=Hartig, G. |author3=Grath, S. |author4=Pohl, H. |author5=Lehmann, J. |author6=Tafer, H. |author7=Donath, A. |author8=Krauss, V. |author9=Eisenhardt, C. |author10=Hertel, J. |author11=Petersen, M. |author12=Mayer, C. |author13=Meusemann, K. |author14=Peters, R.S. |author15=Stadler, P.F. |author16=Beutel, R.G. |author17=Bornberg-Bauer, E. |author18=McKenna, D.D. |author19=Misof, B. | year=2012 |title=Genomic and Morphological Evidence Converge to Resolve the Enigma of Strepsiptera | journal=Current Biology | volume=22 | issue=14 | pages=1309–1313 |doi=10.1016/j.cub.2012.05.018 | pmid=22704986|doi-access=free |bibcode=2012CBio...22.1309N }}</ref><ref name="ColganZhao2014">{{cite journal |last1=Colgan |first1=Donald J. |last2=Zhao |first2=Chenjing |last3=Liu |first3=Xingyue |last4=Yang |first4=Ding |title=Wing Base Structural Data Support the Sister Relationship of Megaloptera and Neuroptera (Insecta: Neuropterida) |journal=PLOS ONE |volume=9 |issue=12 |year=2014 |pages=e114695 |doi=10.1371/journal.pone.0114695|pmid=25502404 |pmc=4263614 |bibcode=2014PLoSO...9k4695Z |doi-access=free }}</ref> groups containing parasitoids are shown in boldface<!--used redundantly alongside bracketed numbers-->, e.g. '''Coleoptera''', with the number of times parasitoidism evolved in the group in parentheses, e.g. '''(10 clades)'''. The approximate number (estimates can vary widely) of parasitoid species<ref name=Mills2009/> out of the total is shown in square brackets, e.g. [2,500 of 400,000].
{{clade
|label1=[[Endopterygota]]
|1={{clade
   |1={{clade
      |label1=[[Neuropterida]]
      |1={{clade
         |1=''[[Raphidioptera]]''
         |2={{clade
            |1=''[[Megaloptera]]''
            |2='''[[Neuroptera]]''' (net-winged insects) '''(1 clade)''' [c. 15 of 6,000]
            }}
         }}
      |label2=[[Coleopterida]]
      |2={{clade
         |1='''[[Coleoptera]]''' (beetles) '''(10 clades)''' [c. 2,500 of 400,000] [[File:Ripiphorid larva on wing of braconid wasp.jpg|alt=[[Ripiphoridae|Ripiphorid]] beetle [[Planidium|triungulin]] larva on the wing of a [[Braconidae|braconid]] wasp|90px]]
         |label2='''(1 clade)'''
         |2='''[[Strepsiptera]]''' (twisted-wing parasites) [600 of 600] [[File:Strepsiptera.png|90px]]
         }}
      }}
   |2={{clade
      |label1=[[Hymenoptera]]
      |1={{clade
         |1=''[[Symphyta]]''
         |label2= '''(1 clade)'''
         |2={{clade
            |1='''[[Orussoidea]]''' (parasitic wood wasps) [75 of 75] [[File:Orussus coronatus.jpg|alt=Parasitic wood wasp|90px]]
            |2='''[[Apocrita]]''' (wasp-waisted insects) [c. 50,000 of 100,000] [[File:Ichneumon_wasp_(Megarhyssa_macrurus_lunato)_(7686081848).jpg|alt=The [[parasitoid wasp]] ''[[Megarhyssa macrurus]]'' ovipositing into host through wood. Her body is c. 50 mm long, her ovipositor c. 100 mm.|90px]]
            }}
         }}
      |label2=[[Panorpida]]
      |2={{clade
         |1={{clade
            |1='''[[Diptera]]''' (true flies) '''(21 clades)''' [c. 17,000 of 125,000] [[File:Stylogaster macalpini (12947561584, cropped) (cropped).jpg|alt=''[[Stylogaster]]'', a [[Conopidae|conopid fly]], showing the long ovipositor|100px]]
             |2={{clade
                |1=''[[Mecoptera]]''
                |2=''[[Siphonaptera]]''
                }}
             }}
          |2={{clade
             |1='''[[Trichoptera]]''' (caddis flies) '''(1 clade)''' [c. 10 of 14,500]
             |2='''[[Lepidoptera]]''' (butterflies, moths) '''(2 clades)''' [c. 40 of 180,000] [[File:Epiricania hagoromo on Euricania facialis (cropped).JPG|alt=Moth ''Epiricania hagoromo'' ([[Epipyropidae]]) feeding on planthopper ''Euricania facialis''|120px]]
             }}
         }}
      }}
   }}
}}

===Hymenoptera===
{{main|Parasitoid wasp}}
[[File:Potter Wasp building mud nest near completion.JPG|thumb|[[Potter wasp]], an idiobiont, building a mud nest; she will [[mass provisioning|provision it]] with paralysed insects, on which she will lay her eggs; she will then seal the nest and provide no further care for her young]]

Within the Hymenoptera, parasitoidism evolved just once, and the many described{{efn|There may be far more species of parasitoid wasp not yet described.}} species of [[parasitoid wasp]]s<ref>{{cite book |title=Hymenoptera of the world : an identification guide to families |date=1993 |publisher=Centre for Land and Biological Resources Research |author1=Goulet, Henri |author2=Huber, John Theodore |isbn=978-0-660-14933-2 |oclc=28024976}}</ref> represent the great majority of species in the order, barring those like the [[ant]]s, [[bee]]s, and [[Vespidae]] wasps that have secondarily lost the parasitoid habit. The parasitoid wasps include some 25,000 [[Ichneumonoidea]], 22,000 [[Chalcidoidea]], 5,500 [[Vespoidea]], 4,000 [[Platygastroidea]], 3,000 [[Chrysidoidea]], 2,300 [[Cynipoidea]], and many smaller families.<ref name=Mills2009/> These often have remarkable life cycles.<ref>{{cite journal |last=Hochberg |first=M. |author2=Elmes, G. W. |author3=Thomas, J. A. |author4=Clarke, R. T. |title=Mechanisms of local persistence in coupled host-parasitoid associations: the case model of Maculinea rebeli and Ichneumon eumerus|year=1996 |volume=351 |issue=1348 |pages=1713–1724 |doi=10.1098/rstb.1996.0153 |journal=Philosophical Transactions of the Royal Society B: Biological Sciences|bibcode=1996RSPTB.351.1713H }}</ref>
They can be classified as either endoparasitic or ectoparasitic according to where they lay their eggs.<ref name="Kapranas2012">{{cite journal |author1=Apostolos, Kapranas |author2=Tena, Alejandro |author3=Luck, Robert F. |title=Dynamic virulence in a parasitoid wasp: the influence of clutch size and sequential oviposition on egg encapsulation |journal=Animal Behaviour |volume=83 |issue=3 |year=2012 |pages=833–838 |doi=10.1016/j.anbehav.2012.01.004|s2cid=54275511 }}</ref> Endoparasitic wasps insert their eggs inside their host, usually as koinobionts, allowing the host to continue to grow (thus providing more food to the wasp larvae), moult, and evade predators. Ectoparasitic wasps deposit theirs outside the host's body, usually as idiobionts, immediately paralysing the host to prevent it from escaping or throwing off the parasite. They often carry the host to a nest where it will remain undisturbed for the wasp larva to feed on.<ref name=Godfray1994/> Most species of wasps attack the eggs or larvae of their host, but some attack adults. [[Oviposition]] depends on finding the host and on evading host defences; the ovipositor is a tube-like organ used to inject eggs into hosts, sometimes much longer than the wasp's body.<ref>{{cite book |author1=Gullan, P. J. |author2=Cranston, P. S. |date=2010 |title=The Insects: An Outline of Entomology |url=https://archive.org/details/insectsoutlineen00pjgu |url-access=limited |publisher=Wiley |edition=4th |pages=[https://archive.org/details/insectsoutlineen00pjgu/page/n388 364], 367 |isbn=978-1-118-84615-5}}</ref><ref name="Gomez, Jose-Maria 2011">{{cite journal | last1=Gomez | first1=Jose-Maria | last2=van Achterberg | first2=Cornelius | year=2011 | title=Oviposition behaviour of four ant parasitoids (Hymenoptera, Braconidae, Euphorinae, Neoneurini and Ichneumonidae, Hybrizontinae), with the description of three new European species | journal=ZooKeys | issue=125 | pages=59–106 | doi=10.3897/zookeys.125.1754| pmid=21998538 | pmc=3185369 | doi-access=free | bibcode=2011ZooK..125U..59V }}</ref><ref name="Quicke Fitton pp. 99–103">{{cite journal | last1=Quicke | first1=D. L. J. | last2=Fitton | first2=M. G. | title=Ovipositor Steering Mechanisms in Parasitic Wasps of the Families Gasteruptiidae and Aulacidae (Hymenoptera) | journal=Proceedings of the Royal Society B: Biological Sciences | publisher=The Royal Society | volume=261 | issue=1360 | date=22 July 1995 | doi=10.1098/rspb.1995.0122 | pages=99–103 | bibcode=1995RSPSB.261...99Q | s2cid=84043987 |quote=The length of the ovipositor compared with the body of the parasitic wasp varies enormously between taxa, from being a fraction of the length of the metasoma to more than 14 times longer than the head and body. (Townes 1975; Achterberg 1986; Compton & Nefdt 1988).}}</ref> Hosts such as ants often behave as if aware of the wasps' presence, making violent movements to prevent oviposition. Wasps may wait for the host to stop moving, and then attack suddenly.<ref name="Achterberg C 1993">Van Achterberg Cornelius; Argaman Q. "Kollasmosoma gen. nov. and a key to the genera of the subfamily Neoneurinae (Hymenoptera: Braconidae)". Zoologische Mededelingen Leiden. 67. (1993):63-74.</ref>

Parasitoid wasps face a range of obstacles to oviposition,<ref name=Godfray1994/> including behavioural, morphological, physiological and immunological defences of their hosts.<ref name="Kapranas2012"/><ref>{{cite journal | last1=Schmidt | first1=O. | last2=Theopold | first2=U. | last3=Strand | first3=M.R. | year=2001 | title=Innate immunity and evasion by insect parasitoids | journal=BioEssays | volume=23 | issue=4| pages=344–351 | doi=10.1002/bies.1049| pmid=11268040 | s2cid=20850885 }}</ref> To thwart this, some wasps inundate their host with their eggs so as to overload its immune system's ability to encapsulate foreign bodies;<ref>{{cite journal |last1=Salt |first1=George | year=1968 | title=The resistance of insect parasitoids to the defense reactions of their hosts | journal=Biological Reviews of the Cambridge Philosophical Society | volume=43 | issue=2| pages=200–232 | doi=10.1111/j.1469-185x.1968.tb00959.x|pmid=4869949 |s2cid=21251615 }}</ref> others introduce a [[Polydnavirus|virus]] which interferes with the host's immune system.<ref>{{cite journal | last1=Summers | first1=M. D. | last2=Dib-Hajj | year=1995 | title=Polydnavirus-facilitated endoparasite protection against host immune defenses | journal=PNAS | volume=92 | issue=1 | pages=29–36 | doi=10.1073/pnas.92.1.29| pmid=7816835 | pmc=42812 | bibcode=1995PNAS...92...29S | doi-access=free }}</ref>
Some parasitoid wasps locate hosts by detecting the chemicals that plants release to defend against insect herbivores.<ref name="Kessler, Andre 2002">{{cite journal | last1=Kessler | first1=Andre | last2=Baldwin | first2=Ian T. | year=2002 | title=Plant Responses to Insect Herbivory: The Emerging Molecular Analysis | journal= Annual Review of Plant Biology  | volume=53 | pages=299–328 | doi=10.1146/annurev.arplant.53.100301.135207| pmid=12221978 }}</ref>

===Other orders===
[[File:Odynerus spinipes^ Vespidae. See parasite note - Flickr - gailhampshire.jpg|thumb|upright|The head of a sessile female [[strepsiptera]]n protruding (lower right) from the abdomen of its wasp host; the male (not shown) has wings]]

The true flies ([[Diptera]]) include several families of parasitoids, the largest of which is the [[Tachinidae]] (some 9,200 species<ref name=Mills2009/>), followed by the [[Bombyliidae]] (some 4,500 species<ref name=Mills2009/>), along with the [[Pipunculidae]] and the [[Conopidae]], which includes parasitoidal genera such as ''[[Stylogaster]]''.  Other families of flies include some [[protelean]] species.<ref name=UWM>{{cite web |url=http://www.entomology.wisc.edu/mbcn/fea506.html |title=Midwest Biological Control News |publisher=Department of Entomology at the University of Wisconsin–Madison |access-date=19 February 2018 |archive-date=5 October 2017 |archive-url=https://web.archive.org/web/20171005235859/http://www.entomology.wisc.edu/mbcn/fea506.html |url-status=dead }}</ref> Some [[Phoridae]] are parasitoids of ants.<ref>{{cite journal |last1=Wuellner |first1=C. T.|display-authors=etal|title=Phorid Fly (Diptera: Phoridae) Oviposition Behavior and Fire Ant (Hymenoptera: Formicidae) Reaction to Attack Differ According to Phorid Species |journal=Annals of the Entomological Society of America |date=2002 |volume=95 |issue=2 |pages=257–266 |doi=10.1603/0013-8746(2002)095[0257:pfdpob]2.0.co;2 |doi-access=free}}</ref><ref>{{Cite journal|last1=Feener |first1=Donald H. Jr.|last2=Jacobs |first2=Lucia F.|last3=Schmidt |first3=Justin O. |date=January 1996 |title=Specialized parasitoid attracted to a pheromone of ants |journal=Animal Behaviour |volume=51 |issue=1 |pages=61–66 |doi=10.1006/anbe.1996.0005|s2cid=16627717|issn=0003-3472}}</ref> Some [[Sarcophagidae|flesh flies]] are parasitoids: for instance ''Emblemasoma auditrix'' is parasitoidal on cicadas, locating its host by sound.<ref>{{cite journal |author1=Köhler, U. |author2=Lakes-Harlan, R. |title=Auditory behaviour of a parasitoid fly (Emblemasoma auditrix, Sarcophagidae, Diptera) |journal=J Comp Physiol A |date=October 2001 |volume=187 |issue=8 |pages=581–587|doi=10.1007/s003590100230 |pmid=11763956 |s2cid=23343345 }}</ref>

The [[Strepsiptera]] (twisted-wing parasites) consist entirely of parasitoids; they usually sterilise their hosts.<ref>{{cite book |author=Whiting, M. F. |editor1=Resh, V. H. |editor2=Cardé, R. T. |year=2003 |title=Encyclopedia of Insects |url=https://archive.org/details/encyclopediainse00resh |url-access=limited |publisher=Academic Press |chapter=Strepsiptera |pages=[https://archive.org/details/encyclopediainse00resh/page/n1122 1094]–1096|isbn=9780125869904 }}</ref>

Two [[Coleoptera|beetle]] families, [[Ripiphoridae]] (450 species<ref name=Mills2009/>)<ref name=ambeetles>{{cite book |author=Falin, Z. H. |year=2002 |chapter=102. Ripiphoridae. Gemminger and Harold 1870 (1853) |pages=431–444 |editor1=Arnett, R.H. Jr |editor2=Thomas, M. C. |editor3=Skelley, P. E. |editor4=Frank, J. H. |title=American beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea |publisher=CRC Press |isbn=978-0-8493-0954-0 |doi=10.1201/9781420041231.ch6|doi-broken-date=12 November 2024 }}</ref><ref>{{cite book |author1=Lawrence, J. F. |author2=Falin, Z. H. |author3=Ślipiński, A. |year=2010 |chapter=Ripiphoridae Gemminger and Harold, 1870 (Gerstaecker, 1855) |pages=538–548 |editor=Leschen, R. A. B. |editor2=Beutel, R. G. |editor3=Lawrence, J. F. |title=Coleoptera, beetles. Volume 2: Morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim) |place=New York |publisher=Walter de Gruyter |isbn=978-3-11-019075-5 |doi=10.1515/9783110911213.538}}</ref> and [[Rhipiceridae]], are largely parasitoids, as are ''[[Aleochara]]'' [[Staphylinidae]]; in all, some 400 staphylinids are parasitoidal.<ref name=Mills2009/><ref name=UWM/><ref name="YamamotoMaruyama2016">{{cite journal |last1=Yamamoto |first1=Shûhei |last2=Maruyama |first2=Munetoshi |title=Revision of the subgenus ''Aleochara'' Gravenhorst of the parasitoid rove beetle genus ''Aleochara'' Gravenhorst of Japan (Coleoptera: Staphylinidae: Aleocharinae) |journal=Zootaxa |volume=4101 |issue=1 |pages=1–68 |year=2016 |doi=10.11646/zootaxa.4101.1.1|pmid=27394607 }}</ref> Some 1,600 species of the large and mainly freeliving family [[Carabidae]] are parasitoids.<ref name=Mills2009/>

A few Neuroptera are parasitoidal; they have larvae that actively search for hosts.<ref>{{cite book |last=Godfray |first=H. C. J. |title=Parasitoids: Behavioral and Evolutionary Ecology |url=https://archive.org/details/parasitoidsbehav0000godf |url-access=registration |year=1994|publisher=Princeton University Press|isbn=978-0-691-00047-3 |page=[https://archive.org/details/parasitoidsbehav0000godf/page/40 40]}}</ref> The larvae of some [[Mantispidae]], subfamily Symphrasinae, are parasitoids of other arthropods including bees and wasps.<ref name=Mills2009>{{cite encyclopedia |author=Mills, N. |date=2009 |edition=2nd |title=Parasitoids |encyclopedia=Encyclopedia of Insects |editor=V. H. Resh |editor2=R. T. Cardé |pages=748–750 |publisher=Elsevier |isbn=978-0123741448 |url=https://books.google.com/books?id=Jk0Hym1yF0cC&pg=PA749}}</ref>

Although nearly all Lepidoptera (butterflies and moths) are herbivorous, a few species are parasitic. The larvae of [[Epipyropidae]] feed on [[Homoptera]] such as leafhoppers and cicadas, and sometimes on other Lepidoptera. The larvae of [[Cyclotornidae]] parasitise first Homoptera and later [[ant]] brood.<ref>{{cite journal |last1=Pierce |first1=Naomi E. |title=Predatory and Parasitic Lepidoptera: Carnivores Living on Plants |journal=Journal of the Lepidopterists' Society |date=1995 |volume=49 |issue=4 |pages=412–453 |url=http://images.peabody.yale.edu/lepsoc/jls/1990s/1995/1995-49(4)412-Pierce.pdf}}</ref> The [[Pyralidae|pyralid moth]] ''[[Chalcoela]]'' has been used in biological control of the wasp ''[[Polistes]]'' in the [[Galapagos Islands]].<ref name="Foottit 2017">{{cite book |last=Foottit |first=Robert G. |title=Insect Biodiversity: Science and Society |url=https://books.google.com/books?id=V3ItDwAAQBAJ&pg=PA606 |year=2017 |publisher=John Wiley & Sons |isbn=978-1-118-94553-7 |page=606}}</ref>

Parasitism is rare in the Trichoptera (caddisflies), but it is found among the [[Hydroptilidae]] (purse-case caddisflies), probably including all 10 species in the ''Orthotrichia aberrans'' group; they parasitise the pupae of other trichopterans.<ref name="Wells 2005">{{cite journal | last=Wells | first=Alice | title=Parasitism by hydroptilid caddisflies (Trichoptera) and seven new species of Hydroptilidae from northern Queensland | journal=Australian Journal of Entomology | volume=44 | issue=4 | year=2005 | doi=10.1111/j.1440-6055.2005.00492.x | pages=385–391}}</ref>

Mites of the family [[Acarophenacidae]] are ectoparasitoids of insect eggs. Unlike the insect parasitoids, it is the adult stage in Acarophenacidae that acts as a parasitoid. Specifically, adult female mites feed on insect eggs and their body swells up with offspring, which eventually emerge as adults.<ref>{{Citation |title=Acarophenacidae |date=2003-01-23 |work=Mites (Acari) for Pest Control |pages=74–77 |editor-last=Gerson |editor-first=Uri |url=https://onlinelibrary.wiley.com/doi/10.1002/9780470750995.ch5 |access-date=2024-08-28 |edition=1 |publisher=Wiley |language=en |doi=10.1002/9780470750995.ch5 |isbn=978-0-632-05658-3 |editor2-last=Smiley |editor2-first=Robert L. |editor3-last=Ochoa |editor3-first=Ronald|url-access=subscription }}</ref>

=== Entomopathogenic fungi ===
All known fungi in the genera ''[[Cordyceps]]'' and ''[[Ophiocordyceps]]'' are endoparasitic.<ref>{{Cite journal |last1=Qu |first1=Shuai-Ling |last2=Li |first2=Su-Su |last3=Li |first3=Dong |last4=Zhao |first4=Pei-Ji |date=2022-07-24 |title=Metabolites and Their Bioactivities from the Genus Cordyceps |journal=Microorganisms |volume=10 |issue=8 |pages=1489 |doi=10.3390/microorganisms10081489 |issn=2076-2607 |pmc=9330831 |pmid=35893547|doi-access=free }}</ref> One of the most notable fungal parasitoids is ''[[Ophiocordyceps unilateralis|O. unilateralis]]'' which infects carpenter ants by breaching the ant's exoskeletons via their spores and growing in the ant's hemocoel as free living yeast cells. Eventually the yeast cells progress to producing nerve toxins to alter the behaviour of the ant causing it to climb and bite onto vegetation, known as the 'death bite'.<ref>{{Cite journal |last1=Evans |first1=Harry C. |last2=Elliot |first2=Simon L. |last3=Hughes |first3=David P. |date=September 1, 2011 |title=''Ophiocordyceps unilateralis'': A keystone species for unraveling ecosystem functioning and biodiversity of fungi in tropical forests? |journal= [[Communicative & Integrative Biology]] |volume=4 |issue=5 |pages=598–602 |doi=10.4161/cib.16721 |pmid=22046474 |pmc=3204140 }}</ref> This approach is so fine-tuned, it causes the ant to bite down on the adaxial leaf midrib, which is the part of the leaf most optimal for the fungus to fruit. In fact, it has been found that in specific circumstances, the time of the death bite is synchronised to solar noon.<ref>{{Cite journal |last1=Hughes |first1=David P |last2=Andersen |first2=Sandra B |last3=Hywel-Jones |first3=Nigel L |last4=Himaman |first4=Winanda |last5=Billen |first5=Johan |last6=Boomsma |first6=Jacobus J |date=2011 |title=Behavioral mechanisms and morphological symptoms of zombie ants dying from fungal infection |journal= [[BMC Ecology]] |language=en |volume=11 |issue=1 |pages=13 |doi=10.1186/1472-6785-11-13 |issn=1472-6785 |pmc=3118224 |pmid=21554670 |doi-access=free |bibcode=2011BMCE...11...13H }}</ref> As much as 40% of the ant's biomass is fungal hyphae at the moment of the death bite.<ref>{{Cite book |last=Sheldrake |first=Merlin |title=Entangled Life |publisher=Vintage |year=2021 |isbn=9781784708276 |pages=107–119 |language=En}}</ref> After the ant dies, the fungus produces a large stalk, growing from the back of the ant's head<ref>{{Cite journal |last1=Pontoppidan |first1=Maj-Britt |last2=Himaman |first2=Winanda |last3=Hywel-Jones |first3=Nigel L. |last4=Boomsma |first4=Jacobus J. |last5=Hughes |first5=David P. |date=2009-03-12 |editor-last=Dornhaus |editor-first=Anna |title=Graveyards on the Move: The Spatio-Temporal Distribution of Dead Ophiocordyceps-Infected Ants |journal= [[PLOS ONE]] |language=en |volume=4 |issue=3 |pages=e4835 |doi=10.1371/journal.pone.0004835 |issn=1932-6203 |pmc=2652714 |pmid=19279680|doi-access=free |bibcode=2009PLoSO...4.4835P }}</ref> which subsequently releases ascospores. These spores are too large to be wind dispersed and instead fall directly to the ground where they produce secondary spores that infect ants as they walk over them.<ref>{{Cite journal |last1=Pontoppidan |first1=Maj-Britt |last2=Himaman |first2=Winanda |last3=Hywel-Jones |first3=Nigel L. |last4=Boomsma |first4=Jacobus J. |last5=Hughes |first5=David P. |date=2009-03-12 |editor-last=Dornhaus |editor-first=Anna |title=Graveyards on the Move: The Spatio-Temporal Distribution of Dead ''Ophiocordyceps''-Infected Ants |journal= [[PLOS ONE]] |language=en |volume=4 |issue=3 |pages=e4835 |doi=10.1371/journal.pone.0004835 |issn=1932-6203 |pmc=2652714 |pmid=19279680|doi-access=free |bibcode=2009PLoSO...4.4835P }}</ref> ''[[Ophiocordyceps sinesis|O. sinesis]]'' is a parasitoid as well, parasitising ghost moth larvae, killing them within 15-25 days, a similar process to that of ''O. unilateralis''.<ref>{{Cite journal |last1=Zhang |first1=Yongjie |last2=Li |first2=Erwei |last3=Wang |first3=Chengshu |last4=Li |first4=Yuling |last5=Liu |first5=Xingzhong |date=2012-03-01 |title=Ophiocordyceps sinensis, the flagship fungus of China: terminology, life strategy and ecology |journal=Mycology |volume=3 |issue=1 |pages=2–10 |doi=10.1080/21501203.2011.654354 |issn=2150-1203|doi-access=free }}</ref>